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Table 2
Soil physical and chemical properties measure in samples
collected in burnt and unburnt zones, 8 and 20 months after Are. Values with ""indicates
significant differences between burnt and unburnt samples at
A
B
the same sampling time. Bold values indicate significant differences between samplings. Significant differences at P < 0.05, both according with t-test
Months after fire
8
20
8
20
Wildfire

3.2.

Unburnt

Burnt

Unburnt

Burnt

Unburnt

Burnt

Unburnt

Burnt

Colony forming units (CFUs)

Microbial abundance in the different microbial groups studied in unburnt areas was similar, except in the case of filamentous fungi which
were higher in unburnt-A than in B. In general, microbial abundance in
unburnt areas decreased with time, although aerobic bacteria in area A
and filamentous fungi in both areas did not show significant differences
between samplings.
Fire effect on viable and cultivable microbial groups studied produced
a general increment compared with unburnt areas during the whole
study period, although this increment in burnt areas tended to decrease
with time, with the exception of fungal abundance in area A which did not
show significant differences compared with unburnt areas (Fig. 2).

Fungal to bacterial ratio (F:B) calculated based on plate count results

was markedly higher in unburnt-A (0.051 0.016 and 0.072 0.017,8
and 20 months after fire respectively) than in unburnt-B (0.022 0.004
and 0.020 0.002, 8 and 20 months after fire respectively). Fire-induced
changes on F:B were different depending on community studied. F:B ratio
decreased with fire in area A (0.016 0.004 and 0.027 0.003, 8 and 20
months after fire respectively), while it increased markedly inarea B
(0.033 0.011 and 0.068 0.013,8 and 20 months after fire respectively). The F:B ratio in both burnt areas was higher in the second
sampling compared with the first one.
3.3.

Microbial biomass

Microbial biomass carbon (Cmic) in unburnt-A was higher than in

unburnt-B during the whole study period (Fig. 3A). When we compare Cmic
regarding SOC (Cmic/SOC ratio) the differences were diluted and were only
significantly higher in unburnt-A than in B in the first sampling (Fig. 3B).
Cmic fire-induced changes were different depending on plant community studied, it appears to be lower in burnt-A than in unbunrt-A,
whilst, Cmic in burnt-B was higher than in the unburnt, although significant differences were not found (Fig. 3A).
The ratio Cmic/SOC in fire affected areas was lower than in the unburnt ones 8 months after fire. Nevertheless, 20 the same ratio was
reached as in unburnt area values in burnt-A, but not in burnt-B (Fig.
3B).
In general, fungal and bacterial PLFA abundances were higher in unburnt-A than in unburnt-B (Fig. 3C, D)
.

Please cite this article as: Brcenas-Moreno, G., et al., Plant community influence on soil microbial response after a wildfire in Sierra Nevada
National Park (Spain), Sci Total Environ (2016), http://dx.doi.org/10.1016/j.scitotenv.2016.05.013

Fig. 2.

CFU mean values (SE, n = 8) of aerobic bacteria (A), spore formers B), actinobacteria (C) and filamentous fungi (D) measured in both areas, 8 and 20 months after fire. The presence of t letter indicate
significantly higher mean value throughout for the same burnt/unburnt and plant community situation, * indicates the significantly higher mean value when compare burnt and unburnt samples in the same
sampling and plant community (t-test, P < 0.05).

Please cite this article as: Brcenas-Moreno, G., et al., Plant community influence on soil microbial response after a wildfire in Sierra Nevada
National Park (Spain), Sci Total Environ (2016), http://dx.doi.org/10.1016/j.scitotenv.2016.05.013

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Fig.3.

Mean values ( SE, n = 8) of soil microbial biomass carbon (Cmic) (A), Cmic/SOC ratio (BD), bacterial PLFA abundance (C) and fungal PLFA abundance (D) measured in both areas,
8 and 20 months after fire. The presence of t letter indicate significantly higher mean value throughout for the same burnt/unburnt and plant community situation, * indicates the
significantly higher mean value when compare burnt and unburnt samples in the same sampling and plant community (t-test, P < 0.05).

When we analysed fungal and bacterial PLFAs 8 months after fire, we

did not find significant differences between burnt and unburnt areas, independent of plant community studied. Nevertheless, 20 months after
fire, fungal and bacterial PLFAs were lower in burnt-A compared with
unburnt-A, while burnt-B showed higher values than the unburnt one
(Fig. 3C, D).

although 20 months after fire burnt-A had decreased to unburnt-A

values, while burnt-B continued above unburnt-B (Fig. 4B).

Fungal to bacteria ratio (F:B) was calculated based on PLFA abundance as well. PLFA-F:B was similar in unburnt-A (0.123 0.020 and
0.183 0.020, 8 and 20 months respectively) and B (0.116 0.017 and
0.160 0.010, 8 and 20 months respectively). Fire induced a marked
decrease in PLFA-F:B in both studied areas reducing values to 0.067
0.011 and 0.102 0.007, at 8 and 20 months respectively in area A and
to 0.036 0.013 and 0.085 0.022, at 8 and 20 months re spectively in
area B. Similar to that described for F-B ratio based on plate count
method, F:B ratio in burnt areas tended to increase with time, although
in this case the increase with time can be observed in unburnt areas as
well.
3.4.

Microbial activity

Microbial activity of heterotrophic microorganisms estimated by

respiration was higher in unburnt-A than in unburnt-B. Burnt-A showed
lower respiration rate than the unburnt 8 months after fire, although in
the second sampling respiration rate values in burnt-A reached unburntA values. Respiration rates in burnt-B were slightly higher than the
unburnt ones during the whole study (Fig. 4A).
Bacterial growth estimated by leucine (Leu) incorporation was similar
in both unburnt areas. Fire effect on bacterial growth showed a marked
increase compared to the unburnt areas in both plant communities,

Please cite this article as: Barcenas-Moreno, G., et al., Plant community influence on soil microbial response after a wildfire in Sierra Nevada
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3.5.

Microbial community composition

Phospholipid fatty acid pattern was studied by principal components

analysis (PCA) including samples in 3 different blocks. First we include in
the analysis all samples corresponding to different plant community (area
A and B) and fire treatment (burnt and unburnt) (Fig. 5A, B). This
analysis distinguishes several components, but 40% of variability was
explained for the two principal components. PC1 explained 21.99% of
variability and appears to be related to temporal variation suffered by the
samples throughout, especially in samples corresponding to area B with
important weight of actinobacteria-relat- ed PLFAs as 10:Me16b, 10:Me18
and 10:Me e17. PC2 accounted 17.62% of variability and appeared to
separate wildfire-affected area from the unburnt ones, with more marked
weight of area B samples in this separation related to higher relative
abundance of 16;0, cy:17 and cy:19 PLFAs.

The second analysis included only samples from area A (Fig. 5C, D).
PC1 accounted 26.2% variability separating burnt from unburnt soil
samples while PC2 accounted 17.6% of variation, separating mainly burnt
samples between first and second sampling. The scores significantly
related to burnt samples during the first sampling were: a15:0, 16:17c,
cy17:0,18:1 7 and 10Me18, while second sampling appears to be more
related to i17:0, a17:0,18:1 9y cy19:0. Unburnt soil samples were more
related with fungal markers, highlighting 18:2 6,9 and 16:1 5.
Third PCA was played with area B-related samples (Fig. 5E, F), with
29.3% of variability accounted in PC1, which separated mainly unburnt
samples through sampling. PC2 accounted 22.2% of variation
distinguishing between burnt and unburnt samples. Scores show cy17 as
main marked associated with burnt samples during the first sampling,
while i15:0, a15:0, i16:0 and cy19:0 appears to be related with burnt
samples without taking into account sampling time. Unburnt soil samples
are related to fungal indicators 18:26,9, and16:15, a

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Fig. 4.

Mean values ( SE, n = 8) of soil respiration rate (A) and bacterial growth (B), measured in both areas, 8 and 20 months after fire. The presence of t letter indicate significantly higher mean value
throughout for the same burnt/unburnt and plant community situation, * indicates the significantly higher mean value when compare burnt and unburnt samples in the same sampling and plant
community (t-test, P < 0.05).

occurred in area A, but in this case 18:1 7, 18:1 9 and 10Me18

markers appear to be related to unburnt samples during the second
sampling as well.
When individual PLFA abundance was analysed (PLFA nmol g -1 dry
soil), the absence of significant differences between the abundance of
different PLFAs between burnt and unburnt samples from area A was
highlighted. Nevertheless, area B showed several PLFA markers with
significantly higher abundance in burnt than in unburnt samples, i15:0,
a15:0, i16:0,16:1 7c, i17:0, a17;0, cy17:0,18:1 7; 10Me18 and cy19.
4. Discussion
4.1. Factors controlling differences in unburnt sites
The unburnt-A site showed higher SWC, EC, C and N contents com-

pared to the unburnt-B site, independently of the sampling time. These

differences could be related to differences in altitude which led to a higher
intensity erosion process and important differences in organic matter
deposition, humification and leaching as revealed by previous works in
the same area (Sanchez-Maranon et al., 2002). Differences induced by
each plant community composition probably related to different fuel load
and quality could also be an influence. In addition, there was a
diminution in SOC and P in unburnt-B from the 1st sampling to the 2nd
one. This diminution could be related to possible contamination by ash
deposition from close-by burnt areas. Ashes could be transported by wind
after fire and re-distributed in the nearest unburnt areas (Pereira et al.,
2015) causing a slight increment in SOC, DOC, N and P that could be still
appreciated 8 months after fire in unbunrt-B but could be consumed by
microorganisms 20 months after fire, causing the diminution of these
nutrients in unburnt-B throughout.
Microbial abundance (Figs. 2, 3) and activity (Fig. 4) were higher in
the unburnt-A than in the unburnt-B, probably due to the higher nutrient content and more favorable growth conditions related to higher

Please cite this article as: Barcenas-Moreno, G., et al., Plant community influence on soil microbial response after a wildfire in Sierra Nevada
National Park (Spain), Sci Total Environ (2016), http://dx.doi.org/10.1016/j.scitotenv.2016.05.013

temperature and water content (Zak et al., 1994). Microbial community

composition differences in unburnt areas are more difficult to interpret,
since seasonal and altitude differences appear to play an important role.
Unbunrt-B microbial community during first sampling was markedly
different to the unburnt-B during second sampling and to the unbunrt-A
in both samplings. These marked changes were probably related to the
higher altitude of this area, which led to more extreme seasonal
differences. So area B suffers lower temperatures during more time than
area A, and higher solar radiation during summer. Differences in
microbial community composition between unburnt-B during first
sampling are mainly related to the presence of 10 Me PLFAs which are
linked to the presence of actinobacteria group (Fig. 5). The higher relative
abundance of this biomarker in unburnt-B during first sampling was
backed up by the abundance of spore formers and actinobacteria CFU,
which were higher during the first sampling, compared with the second
one in both areas. Both, spore-formers and actinobacteria groups are
characterized by the capacity to survive under unfavorable conditions in a
metabolically inactive form (Nicholson et al., 2000; Flardh and Buttner,
2009, Scherr and Nguyen, 2009). The proximity of thaw in the first
sampling compared with the second one could stimulate a massive growth
of inactive microorganisms such as actinobacteria and spore formers
immediately after snow melting in both areas during the first sampling.
Nevertheless, this massive growth could have been neutralized by the
remaining microbial components in area A (earlier thaw at lower altitude)
but not in B, explaining the higher relative abundance of PLFA biomarkers
related to actinobacteria in unburnt-B, 8 months after fire compared with
the remaining samples shown in the PCA (Fig. 5).
In addition, fungal PLFA abundance increased in both areas from the

first sampling to the second one, probably due to temperature increases

after thaw and higher plant activity which increases root exudates release
and more active decomposition of litter accumulated during winter
(Holland and Coleman, 1987; Bardgett et al., 1999). In spite of both
samplings being collected during spring, marked differences in vegetation
flowering were observed between 1st (May) and 2nd (June) sampling in
area B, corroborating that time since thaw was higher in the second
sampling.
4.2. Factors controlling differences in burnt sites
Wildfire effect was different depending on the area studied. Fire effect on
burnt-A was evident 8 months after fire, but 20 months after fire most of
the parameters studied had reached similar values to the un- burnt-A.
Just EC, available P and bacterial CFU (including actinobacteria) were
still significantly higher than unburnt values 20 months after fire, while
fungal abundance estimated by PLFAs was below unburnt values. These
results indicate the capacity of this ecosystem to recover the unburnt
values of different soil parameters in approximately two years after fire,
although the vegetation growth will need several years to reach the same
development as the unburnt-A. The sensitivity of fungi contrasts with
bacterial proliferation after fire which has been associated with pH and
DOC increases in several studies (Pietikainen and Fritze, 1995; MataixSolera et al., 2002; Badia and Marti, 2003; Ponder et al., 2009; BarcenasMoreno and Baath, 2009, Barcenas-Moreno et al., 2011b). DOC and pH
values in burnt-A 8 months after fire were markedly higher than at the
unburnt one, but the differences were diluted 20 months after fire.
Barcenas-Moreno and Baath (2009) in a laboratory heating study
compared fungal and bacterial growth and observe

Please cite this article as: Barcenas-Moreno, G., et al., Plant community influence on soil microbial response after a wildfire in Sierra Nevada
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Fig.5.

Scores (A, C, E) and loading (B, D, F) plots from PCA performed with PLFAs from both areas (A, B), area A (C, D) and area B (E-F), the open symbol

corresponds to unburnt samples while the filled symbol represents burnt samples. The smallest size of symbol indicates first sampling data and the largest
corresponds to the second one. Mean values SE,

n = 4.

lower fungal recovery compared to bacteria suggesting the intense

bacterial proliferation as another possible explanation for fungal recovery
delay, due to competitive interaction with each other. In our study, viable
and cultivable bacteria and bacterial growth showed a marked increase 8
months after fire, and even 20 months after fire compared with unburnt
conditions (Fig. 2). The absence of total recovery of vegetation together
with a possible competition for resources between fungi and bacteria
could be an explanation for fungal recovery delay when most of the soil
parameters appear to reach the unburnt area values.

Fire effect on area-B was still obvious 20 months after fire in most of
the parameters studied. The wildfire induced an increase in the values of
SOC, DOC, Nk and P in burnt-B compared with the unburnt one, even 20
months after fire. This increase in nutrient could be due to ash
incorporation after fire (Ghodrati et al., 1995; Adriano and Weber, 2001;
Demeyer et al., 2001) mainly when combustion of organicmaterial has
been incomplete and charred litter remains (Raison, 1979). This increase
in nutrient, together with pH diminution could indicate low fire intensity
in this area. Also, pH decreases have been observed in low intensity
heated soil (Fernndez et al., 1997) in contrast with the common pH
increases when higher temperatures are reached. This diminution is
usually caused by colloid dehydration and the consequent lost of the soil
pH buffer capacity. Although we should consider the possibility of a
possible difference in the original pH of the burnt-B with regard to the

unburnt-B, before fire incidence, since there are studies that obtained
lower pH values in the same vegetation community in the National Park
which vary from 5.3 to 6.5 (Simn et al., 1994; Snchez-Maran et al.,
2002).

The increment of nutrient and the absence of change in pH due to low

intensity fire appear to be related to microbial response in area-B, where
different microbial parameters studied were at the same or above the
unburnt area values even 20 months after fire. It should b

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pointed that fungal CFU and fungal PLFAs showed the opposite behavior
to that observed in area A. Fungal CFU in burnt-B was twice as abundant
as in the unburnt one and bacterial CFU showed less marked proliferation after fire than those observed in burnt-A. This different pattern could
be related to soil pH differences in both areas, together with differences in
quantity and quality of soil organic matter, fire-induced changes, and
different vegetation post-fire recovery rate. The original soil pH and the
post-fire pH are important factor controlling microbial response, mainly
when fungal and bacterial response is studied. Low pH pre and post-fire
could have an important transcendence in fungal recovery due to original
inoculums and ecological adaptation capacity to new conditions after fire
(Brcenas-Moreno et al., 2016), mainly related to higher DOC
concentration and the possible existence of recalcitrant and toxic
compounds as showed Brcenas-Moreno et al. (2011a) in a laboratory
heating study.

after fire and could be related to fire-induced changes in soil organic

matter quality that need to be studied in depth.
Acknowledgements
This research was supported by the CICYT co-financed FEDER
project CGL2006-11107-C02-01/BOS. We are grateful for the Sierra
Nevada National Park support during the study. We want to express our
gratitude to Prof. Erland Bth for his support and the Microbial Ecology
area of the Biology Department of Lund University where some of the
analyses were realised during FPU fellowship from the Spanish Ministry
of Education to G.B.-M., and to Frances Young for English revision.
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Please cite this article as: Brcenas-Moreno, G., et al., Plant community influence on soil microbial response after a wildfire in Sierra Nevada
National Park (Spain), Sci Total Environ (2016), http://dx.doi.org/10.1016/j.scitotenv.2016.05.013